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Parents might initially produce more offspring than they might be able to raise. However, when offspring demand exceeds their parents ' rearing capacity, parents might shift care towards the offspring which yield greater fitness returns to achieve their optimal brood size via brood reduction. Such favoritism could rely on offspring signaling traits if these inform parents about offspring quality and hence about the pay-offs of their investment. Here we investigated whether favoritism of blue tit (Cyanistes caeruleus) parents for an offspring signal (i.e., ultraviolet (UV) plumage coloration) varies with brood demand. To test this, we experimentally blocked the UV reflectance of yellow breast feathers in half of the nestlings of each brood, and then we sequentially performed two opposing brood size manipulations to vary nestling demand below or above parental rearing capacity. In reduced broods, nestlings begged overall less intensely and gained more body mass, supporting that parental rearing capacities sufficed to satisfy brood demand. Moreover, in reduced broods, UV-blocked nestlings (i.e., low-quality offspring) were fed and prey-tested more often. Yet, they begged more than control nestlings, suggesting that they were perhaps treated differently by other family members or which they may exploit parental preferences beyond actual need (at least in reduced nests). Parents flexibly shifted their feeding rate and favoritism in response to short-term changes in family size, as there was no parental preference for enlarged broods. Such flexible parental feeding rules may allow parents to gain the upper hand in parent-offspring conflict. However, we did not find evidence that parental favoritism facilitated brood reduction, at least in conditions where demand was temporally enhanced. Blue tit parents may produce more offspring than they are able to raise, and they might therefore preferentially invest in high quality offspring that are most likely to survive. Yet when parents are able to raise all their offspring they might even favor those with lowest survival probabilities to maximize reproductive success. Parents hence benefit from flexible feeding strategies for which they rely on offspring signals (i.e., colorful traits), which inform about offspring quality.

In many species, offspring display conspicuous adult-like colouration already early in life, even though they might be very vulnerable to predation at this stage. Yet, the signalling function of adult-like traits in nestlings has been little explored to date. Here, we investigated whether the yellow breast plumage of blue tit (Cyanistes caeruleus) nestlings shows patterns of condition-dependence and hence signals individual quality, as has been described for adult birds. During three consecutive breeding seasons, we therefore explored the association between nestling body mass and three colour components of the yellow breast plumage (i.e., UV chroma, carotenoid chroma and total brightness), considering both within and among nest effects. Carotenoid chroma was not affected by body mass. However, UV chroma and total brightness showed an among-nest effect of body mass, suggesting that they might signal aspects of genetic quality or parental rearing capacity. Interestingly, we also found a within-nest effect of body mass on total brightness, suggesting that this is a good candidate for a condition-dependent signal within the family. Thus, other family members could rely on brightness to adjust their behavioural strategies, such as feeding behaviour in parents. Our study thus reveals that certain colour components of the yellow breast plumage signal different aspects of offspring quality and suggest that they might have a correlated signalling value across life-history stages.

Avian feathers need to be replaced periodically to fulfill their functions, with natural, social, and sexual selection presumably driving the evolution of molting strategies. In temperate birds, a common pattern is to molt feathers immediately after the breeding season, the pre- basic molt. However, some species undergo another molt in winter- spring, the pre-alternate molt. Using a sample of 188 European passerine species, Bayesian phylogenetic mixed models, and correlated evolution analyses, we tested whether the occurrence of the pre-alternate molt was positively associated with prox-ies for sexual selection (sexual selection hypothesis) and nonsexual social selection (social selection hypothesis), and with factors related to feather wear (feather wear hypothesis) and time constraints on the pre- basic molt (time constraints hypothesis). We found that the pre-alternate molt was more frequent in migratory and less gre-garious species inhabiting open/xeric habitats and feeding on the wing, and margin-ally more frequent in species with strong sexual selection and those showing a winter territorial behavior. Moreover, an increase in migratory behavior and sexual selection intensity preceded the acquisition of the pre-alternate molt. These results provide support for the feather wear hypothesis, partial support for the sexual selection and time constraints hypotheses, and no support for the social selection hypothesis.KEYWORDSbirds, feather wear, pre-alternate molt, sexual selection, social selection, time constraintsTAXONOMY CLASSIFICATIONBehavioural ecology; Evolutionary ecology; Life history ecology, European Regional Development Fund; Spanish Ministry of Economy and Competitiveness, Grant/Award Number: CGL2013-48193-C3-1-P, CGL2013-48193-C3-3-P, CGL2017-83103-P, CGL2017-83843-C2-1-P and RYC-2014-15145; Spanish Ministry of Science and Innovation, Grant/Award Number: PID2019-106032GB-I00

During egg laying, females face a trade‐off between self‐maintenance and investment into current reproduction, since providing eggs with resources is energetically demanding, in particular if females lay one egg per day. However, the costs of egg laying not only relate to energetic requirements, but also depend on the availability of specific resources that are vital for egg production and embryonic development. One of these compounds are carotenoids, pigments with immuno‐stimulatory properties, which are crucial during embryonic development. In this study, we explore how carotenoid availability alleviates this trade‐off and facilitates egg laying in a small bird species, the blue tit (Cyanistes caeruleus). Blue tits have among the largest clutch size of all European passerines and they usually lay one egg per day, although laying interruptions are frequent. We performed a lutein supplementation experiment and measured potential consequences for egg laying capacity and egg quality. We found that lutein‐supplemented females had less laying interruptions and thus completed their clutch faster than control females. No effects of treatment were found on the onset of egg laying or clutch size. Experimentally enhanced carotenoid availability did not elevate yolk carotenoid levels or egg mass, but negatively affected eggshell thickness. Our results provide hence evidence on the limiting role of carotenoids during egg laying. However, the benefits of laying faster following lutein supplementation were counterbalanced by a lower accumulation of calcium in the eggshell. Thus, even though single components may constrain egg laying, it is the combined availability of a range of different resources which ultimately determines egg quality and thus embryonic development., This study provides the first evidence of limiting role of carotenoid during egg production. Experimentally enhanced carotenoid availability facilitated egg laying in blue tit females.

In bi-parental species, reproduction is not only a crucial life-history stage where individuals must take ftness-related decisions, but these decisions also need to be adjusted to the behavioural strategies of other individuals. Hence, communication is required, which could be facilitated by informative signals. Yet, these signalling traits might have (co-)evolved in multiple contexts, as various family members usually meet and interact during reproduction. In this study, we experimentally explored for the frst time whether a colourful plumage trait in adults acts as a signal that regulates multiple intra-family interactions in a bird species, the blue tit (Cyanistes caeruleus). We expected that an experimental reduction of adults’ UV/yellow refectance (i.e. a reduction of apparent individual quality) should afect the behavioural strategies of all family members. We found evidence for this at least in adults, since the partners of UV-blocked individuals (either males or females) increased their parental investment — perhaps to compensate for the apparent lower condition of their mates. As the UV-blocked adult did not change its provisioning behaviour, the partner presumably responded to the manipulated signal and not to a behavioural change. However, the ofspring did not co-adjust their begging intensity to the experimental treatment. It is thus possible that they responded to overall parental care rather than the signal. These results suggest that UV/yellow colouration of adult blue tits may act as quality signal revealing the rearing capacity to mates., Open Access funding provided thanks to the CRUE-CSIC agreement with Springer Nature. Grants CGL2016-79390-P and PID2019-106032GB-I00 (to JM) funded by MCIN/AEI/ https://doi.org/ 10.13039/501100011033 and by “ERDF A way of making Europe”. JG-C was supported by Grant BES-2017-079750 and JM by a Ramón y Cajal contract RYC-2014-15145 funded by MCIN/AEI/ https://doi.org/ 10.13039/501100011033 and “ESF Investing in your future”.

In many vertebrates, the enzymatic oxidation of dietary yellow carotenoids generates red keto-carotenoids giving color to ornaments. The oxidase CYP2J19 is here a key effector. Its purported intracellular location suggests a shared biochemical pathway between trait expression and cell functioning. This might guarantee the reliability of red colorations as individual quality signals independent of production costs. We hypothesize that the ornament type (feathers vs. bare parts) and production costs (probably CYP2J19 activity compromising vital functions) could have promoted tissue-specific gene relocation. We review current avian tissue-specific CYP2J19 expression data. Among the ten red-billed species showing CYP2J19 bill expression, only one showed strong hepatic expression. Moreover, a phylogenetically-controlled analysis of 25 red-colored species shows that those producing red bare parts are less likely to have strong hepatic CYP2J19 expression than species with only red plumages. Thus, both production costs and shared pathways might have contributed to the evolution of red signals.

El Reglamento (UE) 2019/1150 nace con la finalidad de proteger a los usuarios profesionales de plataformas de intermediación en línea. La posición privilegiada de estas plataformas permite imponer de forma unilateral las condiciones de su relación contractual. Ante esta situación, desde la Unión Europea se ha decidido aprobar el citado Reglamento, para que las plataformas adecúen sus prácticas comerciales a la buena fe y a la lealtad. De esta forma, se beneficiarán no solo los citados usuarios, sino también los consumidores finales. El Reglamento impone una serie de obligaciones en relación con las condiciones generales contractuales, el uso de datos y un sistema de reclamaciones interno que deben prever estas plataformas. El cumplimiento de estas obligaciones supondrá la adecuación de las plataformas a este nuevo régimen jurídico, razón por la que el Reglamento concede un plazo para ello, hasta el día 12 de julio de 2020, es decir, un poco más de un año desde su aprobación. En el presente artículo se analiza en profundidad los aspectos señalados del Reglamento (UE) 2019/1150 y se estudia la doctrina que se ha pronunciado sobre el mismo.

Factors affecting the evolution of plumage conspicuousness in females are nowadays the focus of debate, and here we explore the possibility that the conspicuousness of female plumage and male participation in nest building are associated in birds. We hypothesize that males that participate in nest building will gain higher fecundity from high-quality ornamented females, whereas ornamented females will adjust fecundity to the costly nest-building behaviour of males. Large-sized species might experience higher costs of nest building and, thus, body size should affect the scenario described above. We used information on male contribution to nest construction (yes/no), male and female conspicuousness (conspicuous or cryptic plumage) and body size of Western Palaearctic passerines. In accordance with the hypothesis, we found that female conspicuousness, in interaction with body mass, was strongly associated with male participation in nest building. For large-sized species, female conspicuousness was positively associated with male participation in nest building. Discrete analyses of correlated evolution rendered evidence of female conspicuousness determining the evolution of male contributions to nest building, with the loss of female conspicuousness occurring more frequently before the loss of male participation in nest building. We discuss possible adaptive scenarios explaining the detected evidence because of mutual sexual selection in males and females, The study was financed by the Ministerio de Economía y Competitividad MINECO (CGL2013-48193-C3-1-P, CGL2013-48193-C3-2-P, CGL2013-48193-C3-3-P and CGL2017-83103-P) and the European Regional Development Fund (FEDER)

El derecho de información reconoce al socio la posibilidad de solicitar informaciones o aclaraciones sobre los asuntos comprendidos en el orden del día de una convocatoria de junta. El ejercicio del derecho, como cualquier otro derecho, no puede ser abusivo, desde un punto de vista objetivo y subjetivo, y debe ser de buena fe. El derecho de información no es absoluto, sino que está sometido a unos límites, pues en determinadas ocasiones los administradores pueden denegar la información, salvo que la petición esté apoyada por un determinado porcentaje de capital. Los motivos que permiten la denegación son los siguientes: a) la información sea innecesaria para la tutela de los derechos del socio; b) que existan razones objetivas para considerar que podrá utilizarse para fines extrasociales; c) que la publicidad de la información perjudique a la sociedad o a sociedades vinculadas. Después de la Ley 31/2014 (por la que se modifica la Ley de Sociedades de Capital para la mejora del gobierno corporativo), la vulneración del derecho de información no legitima al socio para impugnar los acuerdos sociales afectados. La tutela del derecho se logra exigiendo el cumplimiento del derecho y, en su caso, solicitando la indemnización de los daños y perjuicios que la vulneración del derecho haya podido ocasionar.

In bi-parental species, reproduction is not only a crucial life-history stage where individuals must take fitness-relevant decisions, but these decisions also need to be adjusted to the behavioural strategies of a partner. Hence, communication is required, which could be facilitated by condition-dependent signals of parental quality. Yet, these traits have (co-)evolved in multiple contexts within the family, as during reproduction different family members may coincide and interact at the site of breeding. In this study we explore whether a condition-dependent trait acts a quality signal and regulates intra-family interactions in a bird species, the blue tit (Cyanistes caeruleus). As a family is a complex network where signals could be perceived by multiple receivers, we expected that experimentally blocking the reflectance of an adult’s UV/yellow colouration of breast feathers may affect the behavioural strategies of all family members. We found an increase of parental investment in nests with an UV-blocked adult, as the partner compensated for the perceived lower rearing capacity. As the UV-blocked adult did not change its provisioning behaviour, as was to be expected, their partner must have responded to the (manipulated) signal but not to a behavioural change. However, offspring did not co-adjust their begging intensity to a signal of parental quality. Opposite to adults, we propose that offspring respond to the behaviour but not to the parental signal. Overall, our results show experimentally at the first time that UV/yellow colouration of blue tits acts as a quality signal revealing the rearing capacity to mates.

En esta obra se analizan: el contenido del derecho, tanto en un sentido estricto (que es el derecho a realizar preguntas) como en un sentido amplio (que es el llamado derecho documental), que nuestros tribunales han interpretado de forma amplia; los limites del derecho, esto es, que condiciones se deben cumplir para poder ejercitar el derecho y ante que supuestos el organo de administracion podra denegar la entrega de la informacion solicitada; y la tutela del derecho de informacion, analizando la indemnizacion por danos y perjuicios que puede solicitar el socio ante la vulneracion de su derecho y prestando una atencion especial a la impugnacion de los acuerdos sociales, que ha sido objeto de una relevante restriccion

The resolution of family conflicts over parental care involves elaborate behavioral interactions where signals and information exchange play a central role. Usually, the focus is on offspring begging and adult signals and their effect on parental provisioning. Yet, despite offspring of many animal species display structural ornaments during parental dependency, their role in intrafamily conflicts remains practically unexplored. In the blue tit, Cyanistes caeruleus, we experimentally manipulated nestling ultraviolet (UV) color and food availability in 60 broods to investigate if family members flexibly adjust their decisions according to color under different conditions. Feeding rates were not affected by experimental treatments, but plumage color did affect parent–offspring interactions in the form of prey-testings (when a parent places a prey item into a nestling’s gape but removes it again). In nonsupplemented nests, fathers but not mothers tested more prey on UV-reduced offspring, suggesting that fathers evaluate less ornamented chicks when food is scarce. As predicted by theoretical studies, UV-reduced nestlings increased begging in food-supplemented nests, although only to mothers. Moreover, UV-reduced nestlings increased parent-absent begging in all nests, indicating that plumage color affected sib-sib competitive interactions. Finally, UV-reduced offspring gained less body mass and this was probably due to costly intrafamily interactions. Overall, our results suggest that ornamentation during early life plays an important role on social-mediated costs and reveals sex-specific parental strategies according to offspring ornaments., This study was covered by projects CGL2016-79390-P to J.M. and CGL2013-48193-C3-3-P to Juan Moreno (Ministerio de Economía y Competitividad, MINECO, Spain). J.M. is supported by a Ramón y Cajal contract from MINECO.

La convivencia en Derecho español de un régimen de responsabilidad societaria y un régimen de responsabilidad concursal exigible a los administradores de sociedades mercantiles, genera ciertas dudas cuando éstas se encuentran incursas en un procedimiento concursal. En este artículo se van a analizar las diferentes posturas doctrinales y jurisprudencias sobre los efectos del procedimiento concursal en la responsabilidad por deudas ex artículo 367 de la Ley de Sociedades de Capital y en la responsabilidad por daños ex artículos 238 y 241 de la misma Ley. Así mismo, se va a analizar la posible coordinación de estas responsabilidades con la responsabilidad concursal, exigible en supuestos en los que una sociedad mercantil se encuentre en situación de insolvencia.

The offspring of many animals are conspicuous during parental dependence, despite juveniles generally suffering from high predation risk. However, to date, it is unclear whether offspring structural ornaments play a role in intrafamily communication. This is the case of conspicuous plumage in young birds, which is worn unchanged during a long period after fledging, when they still depend on their parents. If plumage color facilitates intrafamily interactions, its role should be more important in large-brooded species, where the strength of intrafamily conflict is potentially stronger. We therefore performed a comparative study in 210 passerine bird species to test whether an offspring structural trait, white plumage, evolves more frequently in lineages with larger clutches. We also explored the number of broods raised per year as another source of intrafamily conflict. First, we found that juvenile whiteness was more frequent in open-nesting species. Moreover, in agreement with our prediction, the presence of juvenile white tail/wing patches was strongly and positively associated with clutch size. This relationship was not due to the strong resemblance between offspring and adult plumage, which was controlled for in the statistical analyses. Moreover, the association remained significant after taking into account predation risk, for which there was information for a subset of species. In contrast, juvenile whiteness was not associated with the number of broods raised per year. These results may suggest that the evolution of juvenile conspicuousness is favored in species with potentially stronger intrabrood sibling conflict., This work was financed by the Ministerio de Economía, Industria y Competitividad MINECO (CGL2016-79390-P, CGL2017‐83103‐P, CGL2017-83843-C2-1-P and CGL2013-48193-C3-1-P) and the European Regional Development Fund (FEDER).

El régimen de impugnación de los acuerdos sociales previsto en la Ley de Sociedades de Capital ha sido objeto de una profunda modificación por la Ley 31/2014, que ha supuesto una restricción del derecho de impugnación. Esta reforma ha supuesto la supresión de la distinción entre acuerdos nulos y anulables. Se crea la categoría de acuerdos impugnables, que ha afectado al catálogo de causas de impugnación y se ha recogido una serie de infracciones ante las que no será posible impugnar un acuerdo social. La reforma también ha afectado al plazo y cómputo de la caducidad de la acción y a diversos aspectos de la legitimación activa, pues además se han unificado estas reglas para todos los acuerdos impugnables con la excepción de los acuerdos contrarios al orden público.

In this essay, the hypothesis that biliverdin pigment plays an antioxidant role in the avian eggshell is proposed. Due to its ability to scavenge free radical species and to reduce mutation, biliverdin potentially counteracts the oxidative action of pathogens that penetrate the eggshell and/or protects the shell membrane from oxidation, thus promoting the proven antioxidant and antimicrobial capacities of the shell membrane itself. Additionally, biliverdin may be able to inhibit viral replication in the eggshell due to its ascribed antiviral properties. Moreover, previous results in other taxa leave open the question of whether biliverdin can be absorbed by the embryo from the eggshell and play a role in embryogenesis. These mechanisms of antioxidant action of eggshell biliverdin remain totally unexplored in birds and in other oviparous animals. The main assumptions and predictions of the antioxidant hypothesis are developed, and directions for future research are proposed., This work was sup-ported by a Ramón y Cajal contract by the Ministerio de Economía, Industria y Competitividad MINECO (RYC-2014-15145) and a research project by the Agencia Estatal de Investigación and the European Regional Development Fund (CGL2016-79390-P/AEI/FEDER, UE).

The objective of this study was to determine Salmonella numbers on retail raw chicken carcasses in Guatemala and to phenotypically characterize the isolates (serotyping and antibiotic susceptibility). In total, 300 chicken carcasses were collected from seven departments in Guatemala. Salmonella numbers were determined using the most-probable-number method following the U. S. Department of Agriculture's Food Safety and Inspection Service protocol. In total, 103 isolates were obtained, all of which were tested for antibiotic susceptibility, whereas 46 isolates were serotyped. Overall, Salmonella prevalence and mean number (mean log most probable number per carcass) was 34.3% and 2.3 (95% confidence interval: 2.1 to 2.5), respectively. Significant differences (P0.05) in Salmonella prevalence were found by storage condition (refrigerated or ambient temperature), market type (wet markets, supermarkets, and independent poultry stores), chicken production system (integrated or nonintegrated production company), and chicken skin color (white or yellow). Chickens produced by integrated companies had lower Salmonella numbers (P0.05) than nonintegrated companies, and white-skin carcasses had lower numbers (P0.05) than yellow-skin carcasses. Among 13 different Salmonella serovars identified, Paratyphi B (34.8%) was most prevalent, followed by Heidelberg (16.3%) and Derby (11.6%). Of all the Salmonella isolates, 59.2% were resistant to one to three antibiotics and 13.6% to four or more antibiotics. Among all the serovars obtained, Salmonella Paratyphi B and Heidelberg were the most resistant to the antibiotics tested. Salmonella levels and antibiotic resistant profiles among isolates from raw poultry at the retail market level were high relative to other reports from North and South America. These data can be used by Guatemalan stakeholders to develop risk assessment models and support further research opportunities to control transmission of Salmonella spp. and antibiotic-resistant isolates from chicken meat to humans.

There is increasing evidence that sexual selection operates in females and not only in males. However, the function of female signals in intrasexual competition has been little studied in species with conventional sex roles. In the Iberian populations of the pied flycatcher (Ficedula hypoleuca), some females express a white forehead patch, a trait that in other European populations, only males exhibit and has become a classical example in studies of sexual selection. Here, we investigated whether the expression of this trait plays a role in female-female competition during early breeding stages. To test this hypothesis, we simulated territorial intrusions by challenging resident females with stuffed female decoys expressing or not a forehead patch. We found that resident females directed more attacks per trial and maintained closer distances to non-patched decoys than to patched ones. Also, patched females were more likely to attack the decoy than non-patched females. Interestingly, females were more aggressive against the decoys when their mate was absent. This may indicate that females relax territory vigilance in the presence of their mate or that males interfere in the interaction between competing females. The behavior of resident males was also observed, although it was not affected by decoy’s patch expression. Our findings suggest that the forehead patch plays a role in female intrasexual competition. If the forehead patch signals fighting ability, as it does in males, we may interpret that non-patched females probably avoided repeating costly agonistic encounters with the most dominant rivals

© 2014 The Authors. Nests are structures built to support and protect eggs and/or offspring from predators, parasites, and adverse weather conditions. Nests are mainly constructed prior to egg laying, meaning that parent birds must make decisions about nest site choice and nest building behavior before the start of egg-laying. Parent birds should be selected to choose nest sites and to build optimally sized nests, yet our current understanding of clutch size-nest size relationships is limited to small-scale studies performed over short time periods. Here, we quantified the relationship between clutch size and nest size, using an exhaustive database of 116 slope estimates based on 17,472 nests of 21 species of hole and non-hole-nesting birds. There was a significant, positive relationship between clutch size and the base area of the nest box or the nest, and this relationship did not differ significantly between open nesting and hole-nesting species. The slope of the relationship showed significant intraspecific and interspecific heterogeneity among four species of secondary hole-nesting species, but also among all 116 slope estimates. The estimated relationship between clutch size and nest box base area in study sites with more than a single size of nest box was not significantly different from the relationship using studies with only a single size of nest box. The slope of the relationship between clutch size and nest base area in different species of birds was significantly negatively related to minimum base area, and less so to maximum base area in a given study. These findings are consistent with the hypothesis that bird species have a general reaction norm reflecting the relationship between nest size and clutch size. Further, they suggest that scientists may influence the clutch size decisions of hole-nesting birds through the provisioning of nest boxes of varying sizes.

Among the cellular pathways conferring protection against oxidative stress, a key role is played by heat-shock proteins, especially HSP70. It has been proposed that blue egg colouration in certain birds, which is related to the deposition of the pigment biliverdin, should be related to basic antioxidant processes in the laying female given the strong antioxidant role of biliverdin. If HSP70 levels express antioxidant defences a higher allocation of the pigment to the clutch could induce a higher response by other components of the antioxidant system like HSP70. In a field study of a breeding population of pied flycatchers Ficedula hypoleuca, a species laying blue-green eggs, we measured spectrometrically the colouration of all eggs and collected blood samples of incubating females one week after clutch completion. We estimated female condition, plasma antioxidant capacity and level of HSP70 in blood cells. HSP70 level was positively related to plasma antioxidant capacity indicating that HSP70 is involved in responses to oxidative stress. Variation in HSP70 level was in good part explained by mean egg blue-green chroma of the clutch when controlling for plasma antioxidant capacity. Results suggest that intensity of eggshell blue colouration in this population is related to a basic system involved in protection against oxidative stress.

Although the role of animal signals in the resolution of family conflicts has been thoroughly studied, it has been typically analysed in isolated two-player interactions. For instance, parents are usually considered as the sole receivers of offspring begging signals or mates the receivers of sexual displays. However, this view does not wholly encompass the dynamic and complex nature of the family scenario. In this essay, we review for the first time the clearest evidence of animal signals found to play a role in more than one family context (e.g. mate–mate, parent–offspring and sib–sib interactions). We then argue that these signals might have coevolved in multiple family contexts because the whole network of related individuals shares genes and similar physiological mechanisms underlying signal expression and perception abilities. Finally, we propose candidate traits that we would expect to function in multiple family contexts and we consider questions that could be addressed in further studies to understand better the evolution of family signals.